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Using Plant Functional Traits to Assess Ecosystem Processes and Community Dynamics in Lowland Fens

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Using Plant Functional Traits to Assess Ecosystem Processes and Community Dynamics in Lowland Fens Using plant functional traits to assess ecosystem processes and community dynamics in lowland fens: understanding the efficacy and applicability of a trait-based approach to plant ecology Fabio Carvalho Gomes da Silva A thesis submitted in partial fulfilment of the requirements of Kingston University London for the award of the degree of Doctor of Philosophy (Ph.D.) April 2017 Abstract The research presented here focuses on the functional aspect of biodiversity of plant communities, with emphasis placed on the functions of species within biological communities and ecosystems, rather than their identity. The prominence of plant functional traits as major contributors to ecosystem functioning is based on underlying mechanisms whereby individual species interact with each other and with their abiotic environment to influence ecological processes on different spatial scales. In this study, particular attention is given to the modulating effects of functional diversity and composition on community dynamics and ecosystem processes (e.g., soil processes relevant to the cycles of carbon and nitrogen), as well as its response to disturbance. A number of green leaf functional traits considered to be pertinent to soil processes and the biogeochemical cycles of carbon and nitrogen were measured from vascular plant species growing in lowland fens in East Anglia, UK. Such habitats are widely recognised as areas of high conservation value for providing numerous benefits to society, including nutrient cycling and soil carbon storage. The set of analyses presented here reveals the implications of different degrees of management intervention for the functional composition of lowland fen plant communities. Overall, the functional diversity of such communities were found to respond strongly to changing disturbance intensity, to significantly interact with abiotic factors to contribute to the provision of ecosystem processes and to exert major effects on species coexistence within plant assemblages. These results confirm the wide applicability of the trait approach when investigating the effects of biodiversity on the stability of biological communities and ecosystems, and is potentially informative to conservation- focused projects that aim to simultaneously enhance biodiversity and the provision of vital ecosystem services. In fact, management intervention was largely found to favour species with a set of traits conducive to enhancing soil carbon storage, lending support to current long term conservation projects that aim to positively influence soil carbon balance. Acknowledgements My unlimited gratitude goes to my First Supervisor Dr Kerry Brown and my co-supervisor Professor Martyn Waller at Kingston University London. Their dedication to this project and constant attention to my work vastly contributed to improving the quality of this thesis and to my experience as a research student at Kingston. I am grateful for the hours of discussions and prompt feedback on any material I produced during the preparation of the individual chapters, even during times of intensive teaching. Particularly, I would like to thank Dr Kerry Brown for his help with the data analysis side of the work and for his positive guidance and attitude through challenging phases of my PhD. Professor Martyn Waller provided expertise in fen systems and was especially helpful in guiding me through the field and lab aspects of the project, including species identification and soil sampling. I also want to thank him for producing the pollen diagrams presented here. I would like to thank Dr Arnoud Boom at The University of Leicester for giving me access to lab facilities and for guiding me through the process of quantifying carbon and nitrogen in soil and plant leaf samples, and for his assistance with methodological issues and the use of stable isotope data. Dr Yit Arn Teh at The University of Aberdeen is also acknowledged for his advice during the initial stages of this work, particularly with soil carbon measurements. I am also grateful to the Norfolk Wildlife Trust (NWT) and Natural England for giving me access to Upton Broad and Woodwalton Fen, respectively. I especially thank Debs Kershaw at the NWT and Alan Bowley at Natural England for providing me with valuable information about the sites and how they are managed, as well as advice on safety during fieldwork. To my PhD colleagues at Kingston University, Gabriel, Penny, Chiara, John and Andreas, for their friendliness and for sharing experiences during my time at Kingston. I want to thank my family, especially my mum and brother, for their long-term support that allowed me to pursue this type of work, and Angie for her invaluable support and positive attitude during the final stages of my work. My gratitude also to the Faculty of Sciences, Engineering and Computing at Kingston University London for the PhD award and for easy access to good quality facilities. I would like to thank the administrative staff at the Faculty who provided support when needed, particularly Rosalind Percival. Table of Contents Abstract Acknowledgements Chapter 1 Introduction 1.1 Biological diversity, community dynamics and ecosystem functioning 1 1.2 The peatland environment.................................................................... 6 1.3 Fen communities................................................................................... 11 Chapter 2 Materials and methods 2.1 Study sites............................................................................................. 15 2.1.1 Upton Fen............................................................................... 15 2.1.2 Woodwalton Fen.................................................................... 18 2.2 Field methods........................................................................................ 21 2.2.1 Vegetation survey................................................................... 21 2.2.2 Vegetation sampling............................................................... 23 2.2.3 Soil sampling........................................................................... 24 2.2.4 Plant litter collection.............................................................. 24 2.2.5 Aboveground biomass (AGB) harvesting................................ 26 2.2.6 Water table height.................................................................. 26 2.3 Laboratory methods.............................................................................. 26 2.3.1 Plant functional traits............................................................. 26 2.3.2 Soil analysis............................................................................. 30 2.3.3 Annual litter production and aboveground biomass estimation......................................................................................... 31 2.4 Data analysis.......................................................................................... 32 2.4.1 Management intensity classification...................................... 32 2.4.2 Functional trait axes............................................................... 32 2.4.3 Community weighted mean (CWM) of plant traits................ 34 Chapter 3 Functional diversity and mean trait composition of lowland fens along a disturbance gradient 3.1 Introduction........................................................................................... 36 3.2 Methods................................................................................................ 41 3.2.1 Functional diversity metrics.................................................... 43 3.2.2 Linear mixed effects models (LMMs)...................................... 45 3.2.3 Fisher’s least significant differences (LSD).............................. 47 3.3 Results................................................................................................... 48 3.3.1 Species richness and functional diversity metrics.................. 48 3.3.2 Community weighted means (CWMs) of plant traits............. 51 3.3.3 Ecosystem processes.............................................................. 53 3.4 Discussion.............................................................................................. 55 3.4.1 Effects of species richness on functional diversity metrics.... 57 3.4.2 Effects of disturbance on taxonomic and functional diversity 59 3.4.3 Effects of disturbance on the CWM of plant traits................. 63 3.4.4 Response of ecosystem processes to disturbance and shifts in plant traits.................................................................................... 65 3.5 Conclusions............................................................................................ 67 Chapter 4 Determining the interactions between environmental variables, plant functional traits and ecosystem processes in lowland fens 4.1 Introduction........................................................................................... 69 4.1.1 Links between ecosystem processes and functioning............ 71 4.1.2 Links between plant functional traits and ecosystem processes.......................................................................................... 72 4.1.3 Hypothesised models linking environmental variables, plant functional traits and ecosystem processes...................................... 74 4.1.3-a Upton and Woodwalton (Model a)......................... 76 4.1.3-b Woodwalton (herb-dominated communities; Model b)..............................................................................
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